Effect of temperature on development, growth and feeding of Coccinella septempunctata and Hippodamia convergens reared on the tobacco aphid, Myzus persicae nicotianae

Preimaginal development, mortality, aphid consumption rate, and size and weight upon reaching the adult stage of the aphidophagous coccinellids Hippodamia convergens Guérin-Méneville and Coccinella septempunctata L. collected from Karditsa, central Greece, were examined at four constant temperatures (14, 17, 20 and 23 °C) and L16:D8. The coccinellids fed on the tobacco aphid, Myzus persicae nicotianae Blackman. Egg, larval and pupal mortality was highest at 14 °C reaching 85.0, 73.8 and 29.4% in H. convergens and 49.3, 75.4 and 58.8% in C. septempunctata, respectively. Total preimaginal development ranged from 57.2 to 70.4 days at 14 °C, and to 16.9 and 22.1 days at 23 °C in H. convergens and C. septempunctata, respectively. Heavier and larger adults of H. convergens were obtained at 17 and 20 ° C. In C. septempunctata temperature did not affect adult weight while the lowest size was observed at 14 and 17 °C. Day-degrees requirements for preimaginal development in H. corvengens were 212.9 above a developmental threshold of 11.0 °C. The corresponding values for C. septempunctata were 281.5 and 10.7 °C. In H. convergens total and daily aphid consumption ranged from 46.8 aphids at 14 °C to 85.0 aphids at 23 °C and from 1.5 aphids at 14 °C to 9.2 aphids at 23 °C, respectively. The corresponding values for C. septempunctata were 112.0 and 2.7 at 14 °C and 157.7 and 12.4 at 23 °C. The results show the high potential of both predators as biological control agents against the tobacco aphid. The knowledge obtained could be essential for their appropriate use and for the improvement of mass rearing systems.     

Spectral reflectance from a broccoli crop with vegetation or soil as background: influence on immigration byBrevicoryne brassicae andMyzus persicae

Light reflectance in five wavebands of the spectrum was measured from broccoli (Brassicae oleracea var.botrytis [L].) interplanted with leguminous cover crops (cover crop background) or broccoli grown as monoculture (bare soil background), and fertilized with compost or synthetic fertilizer. AlateBrevicoryne brassicae (L.) andMyzus persicae (Sulzer) (Homoptera: Aphididae) were monitored in yellow pan water traps and on broccoli leaves. Reflectance intensity was higher with a background of bare soil at all wavebands except blue (400–455 nm) in the early-season. Intensity decreased as broccoli canopy cover increased at all wavebands except blue and green (515–550 nm), declining-most dramatically in the yellow (550–590 nm). Highest late-season intensities were in plots with bare soil background and fertilized with compost (those stressed for nitrogen). Few differences in spectral composition, expressed for each waveband as a percentage of total intensity, were recorded. Numbers of alatae were lowest in cover crop background plots in the early season, reached equivalency with bare soil background by mid-season, and showed highest positive correlations with intensity in the yellow (550–590 nm). Results correspond to laboratory findings that aphids are attracted to higher intensity light, especially in the yellow waveband, and support a phototactic explanation for aphid orientation in the field.     

Oviposition avoidance of parasitized aphid colonies by the syrphid predator Episyrphus balteatus mediated by different cues

Oviposition decisions made by members of a guild of natural enemies can have evolved to avoid intraguild predation, potentially avoiding the disruption of the extraguild prey control. We have studied the oviposition preference of the aphidophagous predator Episyrphus balteatus De Geer (Diptera: Syrphidae) within colonies of Myzus persicae Sulzer (Hemiptera: Aphididae) in the presence of two developmental stages of the aphid parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae). Results from a greenhouse choice experiment showed that E. balteatus females lay significantly fewer eggs in colonies with mummified aphids than in unparasitized colonies. Colonies of parasitized, but not yet mummified did not contain significantly fewer eggs than colonies with unparasitized aphids. In three no-choice experiments, we assessed stimuli coming from aphid honeydew, from the aphids themselves and also from extracts of the aphid bodies, and all of these stimuli mediate the discrimination of mummified aphids from healthy aphids. To a lesser extent these stimuli also contribute to the discrimination against aphids that are parasitized but not yet mummified. These results suggest that the effects of these two species could be complementary for the control of M. persicae, since the species that acts as an intraguild predator, E. balteatus, avoids ovipositing on aphid colonies parasitized by the intraguild prey, A. colemani.     

The effects of azadirachtin-treated diets on the feeding behaviour and fecundity of the peach-potato aphid,Myzus persicae

Adult apterousMyzus persicae, which had fed for 26 h on diet containing azadirachtin at concentrations of 25–100 ppm, produced nymphs at less than half the rate of aphids on control diets. After 50 h on azadirachtin-treated diets nymph production had virtually ceased. After this period, any nymphs produced by adults that had fed on azadirachtin-treated diets were born dead with undeveloped appendages. Honeydew production on azadirachtin-treated diets was unaffected during the first 26 h period but was approximately three times less than on control diets during the subsequent 24 h period.     

Effect of cultivars with varying levels of resistance to aphids on development time,sex ratio,size and longevity of the parasitoid <Emphasis Type="Italic">Aphidius colemani</Emphasis>

The influence of cultivars of common cabbage, Brassica oleracea var. capitata with varying levels of resistance to Brevicoryne brassicae (L.) and Myzus persicae (Sulzer) on key biological characteristics of Aphidius colemani (Viereck) was investigated under laboratory conditions. The total development time for female parasitoids reared on M. persicae did not differ significantly between Minicole (green-leaved, partially resistant with antibiosis factors for B. brassicae) and Derby Day (green-leaved, susceptible to both aphid species); but development was significantly faster (ca 10%) on Ruby Ball (red-leaved, partially resistant with antixenosis factors for B. brassicae). Total development time for females reared on B. brassicae was slightly shorter on Ruby Ball than on Minicole. Males reared on M. persicae developed into adults significantly faster (ca 10%) on Ruby Ball than on Minicole. However, when B. brassicae was the host, no significant variations in development time were observed. Sex ratios, size and longevity of both male and female parasitoids on either host were not significantly influenced by cultivar. The results are discussed in relation to the compatible utilisation of host-plant resistance and biological control in the integrated management of aphids.     

温度对蚜科专化菌暗孢耳霉休眠孢子形成的影响

为探究虫霉休眠孢子形成的关键影响因素,通过孢子浴接种和多浓度生物测定实验,观察蚜科专化菌暗孢耳霉(Conidiobolus obscurus)在5个温度处理(10,15,20,24,28℃)和3个接种浓度梯度下感染寄主桃蚜(Myzus persicae)的情况,并观测不同处理下蚜尸内产休眠孢子的比例。暗孢耳霉对桃蚜的毒力随温度和接种浓度升高而提高。同时,感病致死的蚜尸镜检结果表明:温度显著影响休眠孢子形成,温度越高,形成几率越大;接种浓度的影响次之,在15—24℃间,形成几率随接种浓度提高而增大。这一现象可能的解释:高温环境将使寄主种群增长停滞或消退,暗孢耳霉通过感知环境温度情况,倾向于在较高温时于寄主体内形成休眠孢子来规避接下来可能出现的寄主匮乏期。     

Measurement of aphid feeding rates on artificial diets using 3H-inulin

The radioactivity of the honeydew droplets excreted by young apterous adults of the green peach aphid, Myzus persicae, fed on an artificial diet containing ³H-inulin was a reliable measure of the volume of food ingested by the insects, since almost none of the ingested inulin was absorbed and retained by the insects.

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Zusammenfassung Junge aptere Adulte von Myzus persicae wurden auf künstlicher Diät mit oder ohne Zugabe von radioktiv markiertem Inulin (³H, 100 Ci pro ml; 84.5 cpm pro nl) gehalten. Die Radioaktivität der Blattläuse und ihrer ausgeschiedenen Honigtautropfen wurde in Zeitintervallen nach der Fütterung bestimmt. Die Ergebnisse zeigen, dass Inulin nur in geringer Menge von den Blattläusen absorbiert wird, und, dass die Radioaktivität der ausge schiedenen Honigtautropfen nach Zugabe von ³H-Inulin zur künstlichen Diät ein verlässliches Mass für das Volumen der von ihnen aufgenommenen Nahrung ist.

     

Interactions among the entomopathogenic fungus, Beauveria bassiana (Ascomycota: Hypocreales), the parasitoid, Aphidius matricariae (Hymenoptera: Braconidae), and its host, Myzus persicae (Homoptera: Aphididae)

The effect of the entomopathogenic fungus Beauveria bassiana on the biological characteristics and life table of Aphidius matricariae, a parasitoid of the green peach aphid, Myzus persicae, was studied under laboratory conditions. Aphids were first infected with twice the LC₉₅ of B. bassiana for third-instar M. persicae (2 × 10⁸ conidia/ml). Subsequently, at different intervals they were exposed to 1-day-old mated parasitoid females for 24 h. The number of mummies produced per female and the percentage emergence of the F1 generation differed significantly as a function of the time interval between application of the fungus and exposure to the parasitoid. The interference of B. bassiana on parasitoid development was also studied by first exposing the aphid hosts to the parasitoid for 24 h and subsequently applying B. bassiana. The number of mummies produced by a female A. matricariae varied from 11.8 to 24.8 and was significantly different when the aphids were first exposed to the parasitoids and then treated with B. bassiana 24, 48, 72, and 96 h after exposure. There were no significantly different effects of B. bassiana on net reproductive rate (R₀), mean generation time (T), intrinsic rate (r_m) and the finite rate of increase (λ) of A. matricariae as a result of development in hosts exposed to low or high conidial concentrations (1 × 10², 2 × 10⁸ conidia/ml). The parasitoids developed in infected hosts had lower r_m, λ, T and DT (doubling time) values compared with those that developed in uninfected hosts but no differences were observed in R₀ values. With proper timing, A. matricariae and B. bassiana can be used in combination in the successful biological control of M. persicae.     

Differential thresholds of azadirachtin for feeding deterrence and toxicity in locusts and an aphid

Conclusions The importance of using azadirachtin in the field at levels causing primary antifeedancy may have been overemphasised in the past. Currently, it is recommended that commercial preparations of neem be applied at 50–100 ppm (a.i.). While this would undoubtedly have an antifeedant effect in many phytophagous insects it may also harm beneficial species (Schmutterer, 1990). By lowering the concentration of azadirachtin applied to the crop, food intake by insect pests with low chemoreceptor sensitivity to azadirachtin, e.g. aphids (Nisbet et al., 1993; 1994), would not initially be affected. However, secondary antifeedant effects, IGR and sterilant effects could rapidly manifest themselves and bring about crop protection by reducing insect pest populations without harming natural predator or parasitoid populations.     

Suitability of Aphis fabae, Myzus persicae (Homoptera: Aphididae) and Aleyrodes proletella (Homoptera: Aleyrodidae) as prey for Coccinella undecimpunctata (Coleoptera: Coccinellidae)

The suitability of Aphis fabae Scopoli, Myzus persicae Sulzer and Aleyrodes proletella L. as food sources for Coccinella undecimpunctata L. was evaluated by studying the impact of prey consumption on the predator’s population growth parameters and feeding parameters. Unlike A. proletella, A. fabae and M. persicae supported the development and reproduction of C. undecimpunctata. A. fabae and M. persicae were considered to be essential prey, whereas A. proletella was considered to be an alternative prey. Aphid species showed different degrees of suitability: M. persicae significantly decreased the pre-oviposition period and increased adult longevity, fecundity and fertility compared with A. fabae. Moreover, A. fabae represents a suitable diet for larval development, but is not a suitable food source for adult reproduction. The predator’s population growth parameters, R₀, r_m and λ were increased with M. persicae, whereas T decreased. We found that the 4th instar larvae were the most voracious, particularly when fed on M. persicae; nevertheless, with this prey daily weight gain and feeding efficiency of 4th instar larvae were similar to that of individuals fed with A. fabae.     

Functional significance of the alarm pheromone composition in various morphs of the green peach aphid, Myzus persicae

Three isomers of farnesene occur in the siphon excretions of at least four aphid species: the siphon excretions of Myzus persicae contain (E)--farnesene, as well as (Z,E)--farnesene. The significance of (E)--farnesene as an alarm pheromone was evaluated by the investigation of the ratios and quantities of the above-mentioned farnesene isomers in various morphs of Myzus persicae (Sulz.). It is suggested that a relationship exists between the production of farnesene isomers and the occurrence of morphs.

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Zusammenfassung Es ist bekannt, dass drei Isomere des Farnesens in der Siphonensekret von wenigstens vier Blattläusmorphen vorkommen. So enthielt zum Beispiel das Siphonensekret bei Myzus persicae (E)--, (Z,E)-- und (E,E)--Farnesen. Um die Bedeutung des (E)--Farnesens zu ermitteln, haben wir die Mengen und Verhältnisse der Farnesenisomere in verschiedenen Morphen von Myzus persicae bestimmt. Die Ergebnisse deuten darauf hin, dass eine Beziehung zwischen der Produktion von Farnesenisomeren und der Bildung der Morphe in dieser Blattläus besteht.

     

Mechanisms of host discrimination and intraspecific competition in the aphid parasitoid Ephedrus cerasicola

The aphid parasitoid Ephedrus cerasicola Starý (Hymenoptera: Aphidiidae) discriminates between parasitized and unparasitized aphid hosts (Myzus persicae (Sulzer), Homoptera: Aphididae) in two different ways. By antennal contact with the parasitized host, it can detect an external marking pheromone which is effective only for a few hours. The behaviour of the parasitoid also indicated that it responded to an internal marker, probably by means of chemoreceptors on the ovipositor. The parasitoid behaviour was studied in encounters with parasitized hosts 0–10 min, 2–3 h, 6–7 h and 23–24 h after the first parasitization. Aphids were superparasitized within 1 h or after 24 h, and aphids in each experiment were later dissected at two different times. It is suggested that the supernumerary offspring were eliminated by chemical means, whether the time between the two ovipositions was 0–1 h or 24 h.

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Résumé Le parasitoïde de puceron, E. cerasicola Starý (Hym.: Aphidiidae) sélectionne les pucerons, Myzus persicae (Sulzer), parasités ou non, de deux façons différentes. Il peut percevoir une phéromone externe de marquage perceptible seulement pendant quelques heures par contact antennaire. Son comportement montre qu'il réagit aussi à un marquage interne, probablement par des chémorécepteurs de sa tarière. Le comportement du parasitoïde a été étudié en le mettant en contact avec des hôtes parasités depuis 0–10 min, 2–3 h, 6–7 h et 23–24 h. Les pucerons ont été surparasités pendant la première heure ou après 24 heures. Les pucerons de toutes les expériences ont été disséqués à deux moments différents. Ceci a conduit à estimer que les descendants en surnombre ont été éliminés chimiquement quand la période entre deux pontes successives était inférieure à 1 h ou supérieure à 24 h.

     

Potential of Lecanicillium species for dual microbial control of aphids and the cucumber powdery mildew fungus, Sphaerotheca fuliginea

Detached leaf disc bioassays were conducted against cucumber powdery mildew and three species of aphid with three entomopathogenic species of Lecanicillium; Lecanicillium longisporum (Vertalec^®), Lecanicillium attenuatum (CS625), and an unidentified isolate (DAOM198499). The three Lecanicillium species had high virulence against the aphids Myzus persicae, Macrosiphum euphorbiae and Aulacorthum solani with the exception of DAOM 198499, which demonstrated reduced virulence to A. solani with an LT₅₀ of 6.4 days. Otherwise, LT₅₀ ranged between two and four days. Suspensions of conidia and blastospores of the Lecanicillium species were also applied onto 15 mm leaf discs dissected from cucumber plants previously inoculated with Sphaerotheca fuliginea. Powdery mildew did not develop when the Lecanicillium applications were made one and eight days after S. fuliginea inoculations. When Lecanicillium was applied to highly infected leaf discs 11 and 15 days after S. fuliginea inoculation, the application suppressed subsequent production of S. fuliginea spores as compared to the controls. These results suggest the potential of a dual role for Lecanicillium spp. as microbial control agents against aphids and powdery mildew.     

Patch residence decisions made by Aphidius colemani in the presence of a facultative predator

One of the key questions in ecology is how animals optimally allocate their time in an environment with patchily distributed resources and competing organisms. Here we investigate the effects that an aphid predator, Macrolophus caliginosus (Wagner) (Hemiptera: Miridae), has on the searching behavior and the patch residence decisions of an aphid parasitoid, Aphidius colemani (Viereck) (Hymenoptera: Aphidiidae). A computer programme was designed that allowed the recording and saving of direct observations. The time allocated to different activities by a female parasitoid wasp in the presence or absence of the predator M. caliginosus was investigated. The experiments were conducted under controlled environment conditions using leaves of sweet pepper, Capsicum annuum L. (Solanaceae) and Myzus persicae (Sulzer) (Hemiptera: Aphididae) as the host plant–prey species system. The parasitoid spent significantly less time on ‘secondary’ activities, such as preening and resting, when the predator was present. Survival analysis showed that the parasitoid had a higher patch-leaving tendency when the predator was present.     

Genetic variability in host plant adaptation of the green peach aphid, Myzus persicae

The green peach aphid, Myzus persicae (Sulz.), is polyphagous on over 400 plant species in more than 50 families. Phenotypic plasticity of individuals and genetic variability in the population presumably contribute to this polyphagy. The genetic variability in field populations of M. persicae was assessed with respect to their adaptation to sugar beets and potatoes. An analysis of more than 1 000 clones, sampled during 1980, 1981 and 1982 from different host plants in the field, revealed a wide genetic variability in host plant adaptation to sugar beets as well as to potatoes. Both traits seem to be inherited independently from each other and do not correlate with clone-specific host plant preference of apterous adults. The aphid M. persicae can be characterized as a polyphagous insect species with a wide, continuously distributed variability and a broad phenotypic plasticity. A general differentiation of herbivorous species into generalists and specialists tends to ignore the genetic component in the complex of insect-plant relationships.

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Zusammenfassung Die Grüne Pfirsichblattlaus Myzus persicae (Sulz.) lebt polyphag an über 400 Pflanzenarten in mehr als 50 Pflanzenfamilien. Ein breites Nahrungsspektrum einer Art ergibt sich jedoch aus der phänotypischen Plastizität des Individuums oder einer Klonlinie und der genetischen Variabilität der Population. Felpopulationen der Grünen Pfirsichblattlaus wurden auf ihre genetische Variabilität bezüglich der Wirtspflanzenanpassung an Rübe und Kartoffel untersucht. Eine Analyse von mehr als 1 000 Klonen, die über die Jahre 1980, 1981 und 1982 im Rheinland gesammelt wurden, lassen eine breite Variabilität in der Wirtspflanzenanpassung der Population erkennen. Beide Merkmale scheinen unabhängig voneinander vererbt zu werden und zeigen keine Beziehung zum Wirtswahlverhalten adult apterer Läuse der entsprechenden Klone. Die Art M. persicae kann daher charakterisiert werden als eine polyphage Insektenart mit einer breiten genetischen Variabilität und einer grossen phänotypischen Plastizität. Eine generelle Differenzierung von Herbivoren in Generalisten und Spezialisten vernachlässigt die genetische Komponente in der komplexen Beziehung zwischen Insekten un ihren Wirtspflanzen.

     

The production of different morphs by alate viviparae of the aphid Myzus persicae temporarily exposed to long scotophases

The development of alatae of the green peach aphid, Myzus persicae, as gynoparae rather than as virginoparae was investigated with regard to the number of exposures to a long-night (LN) regime of 15 h darkness per diem which the aphids experienced before and/or after their birth. The minimum number of exposures to LN that resulted in all of the alatae developing into gynoparae was two prenatal plus one postnatal or one prenatal plus two postnatal, provided the scotophases in these treatments were at least 12 h long. A cumulative effect of several successive exposures to LN was also evident when the presumptive alatae were exposed to LN either from birth or not until several days after birth. Fewer exposures to LN were needed in the former case.

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Zusammenfassung Die Entwicklung von Alatae der grünen Pfirsichblattlaus, Myzus persicae, hauptsächlich zu Gynoparae, eher als zu Virginoparae, wurde im Hinblick auf den Einfluss der Anzahl an Langnächten (LN: 15 Studen Dunkelheit pro Tag), denen die Aphiden vor und/oder nach der Geburt ausgesetzt waren, untersucht. Zur ausschliesslichen Entwicklung aller Alatae zu Gynoparae waren mindestens 2 prenatale und eine postnatale LN-Exposition oder eine prenatale und 2 postnatale LN-Expositionen notwendig, vorausgesetzt die Dunkelphasen betrugen mindestens 12 Stunden. Ausserdem zeigte sich ein kumulativer Effect durch mehrere, aufeinanderfolgende LN-Expositionen, wenn die Alatae diesen von Geburt an, oder einige Tage nach der Geburt, ausgesetzt waren. Im ersten Fall waren weniger LN-Expositionen notwendig.

     

Hyperpredation by generalist predatory mites disrupts biological control of aphids by the aphidophagous gall midge Aphidoletes aphidimyza

Biological control of different species of pest with various species of generalist predators can potentially disrupt the control of pests through predator-predator interactions. We evaluate the impact of three species of generalist predatory mites on the biological control of green peach aphids, Myzus persicae (Sulzer) with the aphidophagous gall midge Aphidoletes aphidimyza (Rondani). The predatory mites tested were Neoseiulus cucumeris (Oudemans), Iphiseius degenerans (Berlese) and Amblyseius swirskii Athias–Henriot, which are all commonly used for pest control in greenhouse sweet pepper. All three species of predatory mites were found to feed on eggs of A. aphidimyza, even in the presence of abundant sweet pepper pollen, an alternative food source for the predatory mites. In a greenhouse experiment on sweet pepper, all three predators significantly reduced population densities of A. aphidimyza, but aphid densities only increased significantly in the presence of A. swirskii when compared to the treatment with A. aphidimyza only. This stronger effect of A. swirskii can be explained by the higher population densities that this predator reached on sweet pepper plants compared to the other two predator species. An additional experiment showed that female predatory midges do not avoid oviposition sites with the predator A. swirskii. On the contrary, they even deposited more eggs on plants with predatory mites than on plants without. Hence, this study shows that disruption of aphid control by predatory mites is a realistic scenario in sweet pepper, and needs to be considered when optimizing biological control strategies.     

Host plant changes produced by the aphid Sipha flava: consequences for aphid feeding behaviour and growth

Induced plant responses may affect the behaviour and growth of the attacking herbivore insect. The aphid Sipha flava (Forbes) produces reddish spots on the infested leaf of its host plant Sorghum halepense (L.). In order to assess the consequences on the aphid of this presumptive induced plant response, we studied the feeding behaviour and growth of S. flava on previously infested and non-infested leaves of S. halepense. Considering that the reddish pigment could play a defensive role, its effect on aphid survival was determined in artificial diets. In addition, changes in the histology of the leaf and the chemical nature of the induced pigment were also studied. Aphids devoted a significantly shorter total time to non-penetration activities in infested than in non-infested leaves. Time before the first phloem ingestion tended to be shorter in infested leaves. The mean relative growth rate of S. flava nymphs was significantly higher on infested than on non-infested leaves. Survival of aphids on diet containing the reddish extract was not significantly different from that on the control diet. Infestation of S. halepense by S. flava produced a reddish coloration in the leaf, which was identified as an anthocyanin by UV-visible spectrometry. Light microscopy showed that only mesophyll cells of previously infested plants presented swelled, dispersed, and heterogeneously stained chloroplasts with a higher accumulation of starch granules, no grana arranged in stacks, and reduction in the amount of inner membranes (thylakoids), relatively to chloroplasts of non-infested leaves. Scanning electron micrographs of leaf surface revealed reduced presence of crystalline epicuticular waxes of epidermal cells in infested leaves as compared to non-infested ones. The main conclusion is that the attack of S. flava to S. halepense leaves induced plant susceptibility where aphid feeding behaviour and growth were both enhanced on previously infested leaves.     

Phylogeographic patterns in Leccinum sect. Scabra and the status of the arctic-alpine species L. rotundifoliae

We investigated inter- and intraspecific phylogenetic relationships in the ectomycorrhizal fungal genus Leccinum section Scabra. Species of this section are exclusively associated with Betula and occur throughout the Northern Hemisphere. We compared the phylogenetic relationships of arctic, alpine, boreal and temperate accessions of section Scabra based on DNA sequences of the single-copy nuclear gene Gapdh and the multiple-copy nuclear region 5.8S-ITS2. Exclusively arctic lineages were not detected in species that occur both in arctic-alpine or boreal regions, except in L. rotundifoliae that was restricted to cold climates. L. scabrum and L. holopus showed an intercontinental phylogeographic pattern, and L. variicolor showed a pattern unrelated to geographical distribution. Molecular clock estimates indicated that L. rotundifoliae is as old as other species in section Scabra. Individual gene trees suggest that interspecific hybridisation occurred several times in the evolution of section Scabra.